PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 129
GEOLOGICA CARPATHICA, 55, 2, BRATISLAVA, APRIL 2004
PECTINID BIVALVES FROM THE GRUND FORMATION (LOWER
BADENIAN, MIDDLE MIOCENE, ALPINE-CARPATHIAN FOREDEEP)
TAXONOMIC REVISION AND STRATIGRAPHIC SIGNIFICANCE
Department of Paleontology, University of Vienna, Geozentrum, Althanstrasse 14, A-1090 Vienna, Austria; firstname.lastname@example.org
(Manuscript received June 5, 2003; accepted in revised form December 16, 2003)
Abstract: The present study investigates the pectinid bivalve record characterizing the Middle Miocene Grund Forma-
tion from the Lower Austrian part of the Alpine-Carpathian Foredeep. The results provide the first comprehensive
compilation of that pectinid assemblage, based on the recent excavation material and historic collections. Updating and
revision of genus and species level taxa excluded polyphyletic taxa and allowed a more accurate correlation with occur-
rences from adjacent regions distributed across the NE Atlantic, Mediterranean, Paratethys and Indian Ocean. Hence 10
species level taxa Chlamys trilirata, Hinnites crispus, Crassadoma multistriata, Aequipecten malvinae, Ae. macrotis,
Costellamussiopecten cristatus badense, Oopecten solarium, Macrochlamis nodosiformis, M. tournali and Pecten
subarcuatus styriacus were identified. Thereby the subgenus Costellamussiopecten was separated from Amussiopecten,
elevated to the genus level and applied to the lineage including the European Amusium C. cristatus. Macrochlamis
has been regarded as a valid name, and a senior synonym of Gigantopecten. Chlamys represents a still unavailable
genus-name closely related to the Chlamys group. The species level revisions include discovery of H. crispus based on
previously erroneously identified specimens. Moreover the new material brought the first evidence of the presence of Ae.
malvinae and Ch. trilirata in the Grund Formation. The latter three taxa greatly enhanced the stratigraphic signature of
this pectinid assemblage allowing an accurate inference of the Early to Middle Badenian (regional Central Paratethys
geochronological unit isochronous with the Langhian, that is early Middle Miocene) age and a regional correlation with
the Lower Lagenidae Zone for the NE Austrian Neogene basins.
Key words: Middle Miocene, Lower Badenian, Central Paratethys, Alpine-Carpathian Foredeep, biostratigraphy,
taxonomic revision, pectinid bivalves.
The pectinid bivalves are substantial contributors to the Grund
Formation molluscan fauna (e.g. Kautsky 1928), representing
the oldest Middle Miocene molluscan assemblage of the Al-
pine Molasse Zone in Lower Austria. Those characteristic in-
habitants of the past and recent marine shelf regions went
through a massive radiation event in the Lower Miocene and
rose consequently to typical elements of the Neogene shallow
benthic fauna (Ben Moussa & Demarcq 1992). Due to their
abundance, intensive speciation, common short species dura-
tion but also due to enhanced preservation potential due to
predominantly calcite shell mineralogy (e.g. Taylor et al.
1969; Carter et al. 1998), this group of bivalves represents an
important biostratigraphic tools in shallow marine environ-
ments from Oligocene to recent times. Indeed the Paleogene-
Neogene pectinid biozonations exist both for the Mediterranean
(Demarcq 1990, 1992) as well as for the Paratethyan regions
(Báldi & Sene 1975; Bohn-Havas et al. 1987; Studencka
The aim of the present study is to provide the first compre-
hensive compilation of the pectinid bivalve record contribut-
ing the Grund Formation, based on the published material,
historic collection of the Natural History Museum in Vienna
and the University of Vienna Department of Paleontology
and on the recent excavation material. Updating and revision
of genus and species level taxa should exclude the polyphylet-
ic taxa and allow accurate species level correlations with other
well defined occurrences. This will enable the construction of
a most accurate stratigraphic range chart for the assemblage
providing finally a solid base for analysis of the faunal bios-
Paleogeography, age and regional geological setting
The studied assemblage derives from the Alpine Molasse
Zone situated in the western part of the Alpine-Carpathian
Foredeep (Fig. 1). In the Middle Miocene the region belonged
to the western part of the Paratethys Sea, termed also the Cen-
tral Paratethys. The Western Paratethys Sea that occupied the
Alpine Foredeep westwards from the Bohemian Massif termi-
nated in the latest Ottnangian. The regional geodynamic histo-
ry influenced strongly the Central Paratethys deposition and
environment resulting in difficulties in correlations with the
standard stratigraphic divisions and development of the re-
gional chronostratigraphic division. Hence the Grund Forma-
tion belongs to the regional Badenian Stage corresponding to
the lower part of the Middle Miocene.
The fully marine deposits of the Grund Formation (Fig. 2)
overlay similar deposits of the Karpatian Laa Formation com-
plicating commonly their distinction in the field (Rögl et al.
Fig. 1. Regional geological and paleogeographical position of stud-
ied samples from the Lower Badenian Grund Formation (modified
after Mandic et al. 2002). The Molasse Zone corresponds with the
Alpine-Carpathian Foredeep, whereas the Vienna Basin is an intra
mountainous strike-slip basin raised through extensional move-
ments between the Alpidic and the Carpathian Units. In the Middle
Miocene the region belonged to the western part of the Paratethys
Sea, termed also the Central Paratethys. The Western Paratethys
Sea that occupied the Alpine Foredeep westwards from the Bohe-
mian Massif terminated in the latest Ottnangian (cf. Fig. 3).
2002). The mollusc bearing outcrops of the Grund Formation
are scattered in wine cellars, representing the upper part of the
250 m thick succession. This succession belongs to the NN5
Calcareous Nannoplankton Zone and bears foraminiferal as-
semblage with Praeorbulina glomerosa circularis and Uvige-
rina macrocarinata and in its upper part also Orbulina sutur-
alis (Rögl et al. 2002; Æoriæ et al. 2004). The site at Grund
was artificially outcropped during the excavation seasons in
1998 and 1999 (Zuschin et al. 1999; Roetzel & Pervesler
2004). NN5 datum was inferred by Æoriæ & vábenická
(2004) throughout the section, whereas the introduction of
rare P. glomerosa circularis and Uvigerina macrocarinata
was found by Rögl et al. (2002) in the upper part of the com-
posite section, correlating with the paleoenvironmental deep-
ening. Hence the FAD of the P. glomerosa circularis with
16.0 Ma precede distinctly the base of the NN5 Zone (Rögl et
al. 2002). Orbulina suturalis, possibly requiring still deeper
marine settings, was not found at Grund excavation sites 1998
and 1999. The paleomagnetic survey of the site proved, how-
ever, a normal polarity, corresponding with the Chron
C5Bn.2n (Fig. 3; Æoriæ et al. 2004) having the lower boundary
at 15.1 Ma and correlating exactly with the base of the M6
Zone (defined indeed by the FAD of Orbulina suturalis) by
Berggren et al. (1995). For the latter reasons the mollusc bear-
ing horizons at latter site must succeed the base of the Middle
Miocene by at least 1.3 Ma (cf. Rögl et al. 2002). This site
along with other fossiliferous localities of the Grund Forma-
tion are all attributed previously by Grill (1968) into the Lower
Lagenidae Zone. As argued by Rögl et al. (2002) the cooccur-
rence of Praeorbulina glomerosa circularis and Orbulina su-
turalis is typical for the upper part of the Lower Lagenidae
The studied material derives from different sites in the Grund
Formation (Figs. 1, 2). Except for a very fine surface abrasion
the shells are exceptionally well preserved. The juvenile as
well as adult specimens are abundantly present.
This study is based on the material sampled during the exca-
vation 1998 at Grund (Fig. 2, point 1). The series of ditches ex-
posed a predominantly silty and sandy siliciclastic succession
including numerous specimen rich shell beds of tempestitic or-
igin (Roetzel et al. 1999; Zuschin et al. 2001; Roetzel & Per-
vesler 2004). The complete shell material has apparently been
transported from a shallower subtidal sea bottom area into a
deeper marine environment inhabited mono-specifically by a
chemosymbiont bearing bivalve Thyasira michelottii (Hoernes).
The material is in the collection of the Department of Paleon-
tology, University of Vienna (abbreviated in text as DPUV1).
The specimen richer material derives, however, from a field
ditch E Grund (Fig. 2, point 2) in lateral eastward continuation
to the former locality. Its sediment character the yellowish,
silty fine sand and the taphonomic features of shells is iden-
tical to that of the Grund Excavation 1998 site. The site posi-
tion is on the acre, about 100 m south-southwest wards from
the railroad crossing between Grund and Wullersdorf (Fred
Rögl, pers. com. 2003). Fritz Steininger (Senckenberg Muse-
um Frankfurt) & Fred Rögl (Natural History Museum Vienna)
took a large bulk sample there, that has been sorted out and
stored in the Stratigraphic Collection of the Department of Pa-
leontology, University of Vienna (abbreviated in text as
Additional studied material derives from the Locality and
Systematic Collections of the Department of Geology and Pa-
leontology at the Natural History Museum in Vienna. The lo-
cality specifications of investigated specimens are Grund,
Immendorf, Guntersdorf and Windpassing (Fig. 2, underlined
village names). The exact position of historic localities is un-
known. The systematic collection (abbreviated in text as
NHMs) comprises specimens investigated and documented by
Hörnes (1867) and Kautsky (1928). The locality collection
comprises the material studied by Sieber (1947, 1949) (abbre-
viated in text as NHMls), but also the material derived from
the Grund Excavation 1998 (abbreviated in text as NHMlv).
The latter specimens derive from a large microvertebrate bulk
sample collected by Gudrun Höck & Mathias Harzhauser (both
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 131
Fig. 2. Geographical and geological position of sites from which the
studied pectinids come. The fully marine deposits of the Grund For-
mation overlay similar deposits of the Karpatian Laa Formation gen-
erally complicating their distinction in the field. Biogenic limestone
of the Badenian Mailberg Formation intercalates with the topmost
part of the Grund Formation. The Sarmatian sands in the marginal,
restricted marine facies transgressively overlay later deposits and
are exposed in the southeastern part of the region, around Holla-
brunn. Conglomerates of the Pannonian Hollabrunn-Mistelbach For-
mation on the top of the succession represent an ancient EW strik-
ing fluvial stream. The indicated sites are: (1) Excavation Grund
1998 and (2) Field ditch E Grund (cf. text). The underlined names of
villages are site specifications of the studied material from the Natu-
ral History Museum Collections. The more precise positions of the
historical sites are however unknown (illustration combines geology
by Roetzel & Pervesler (2004) and geography by the Austrian Map
1:200,000, BEV, Vienna).
Natural History Museum Vienna). The sample derives from
the basal bed of the section B1 of Zuschin et al. (2001) and
corresponds to the sample specification GRU B1-1 of Roetzel
& Pervesler (2004). It represents a composite shell bed with a
silty matrix in the lower 20 cm and a coarse sandy matrix in
the upper 10 cm. The base of the upper part is characterized by
the occurrence of flattened mud clasts. Beside predominantly
fully marine molluscs, also land gastropods, brackish bivalves
and remains of marine and terrestrial vertebrates are present.
Results and discussions
Faunal composition and taxonomic revisions
The investigated samples documents ten different pectinid
species contributing to the Grund Formation. These are:
1. Chlamys trilirata
2. Hinnites crispus
3. Crassadoma multistriata
4. Aequipecten malvinae
5. Aequipecten macrotis
6. Costellamussiopecten cristatus badense
7. Oopecten solarium
8. Macrochlamis nodosiformis
9. Macrochlamis tournali
10. Pecten subarcuatus styriacus
Three species among them can be considered new for the
Grund Formation. This is an unexpected result as the taxo-
nomic content of the famous assemblage was already the sub-
ject of numerous previous studies (e.g. Hörnes 1867; Kautsky
1928; Sieber 1947, 1949; Schultz 2001). In particular, where-
as Aequipecten malvinae and Hinnites cristatus represent
completely new records, a note on the presence of Chlamys
trilirata was provided recently by Mandic & Harzhauser
(2003). The specimens of the latter species were recently ex-
tracted from samples from the field ditch E Grund and from
the Grund excavation 1998 material. Correspondingly
Aequipecten malvinae derived also from the Grund excavation
and from the field ditch E Grund. Both species were indeed
absent from the historical collections. Hence only the speci-
mens of Hinnites cristatus were found within the material al-
ready investigated by Hörnes (1867) and Schultz (2001). As
discussed below these represent the erroneous identifications
of Hinnites brussoni leufroyi.
Among the new Grund material however no traces of Macro-
chlamis tournali, M. nodosiformis, Oopecten solarium, Cos-
tellamussiopecten cristatus badense and Hinnites crispus
were found. Thereby whereas Pecten subarcuatus styriacus,
Aequipecten macrotis and Crassadoma multistriata are rela-
tively common contributors, the two newly found species are
rather rare. Finally, it can be stated that the new fauna includes
many juveniles, but lacks the larger individuals, which is an
apparent sorting effect (cf. Zuschin et al. submitted).
Two taxa mentioned by previous workers are not included
in the present study. These are Aequipecten scabrellus an d
Oppenheimopecten aduncus (cf. Schultz 2001). The first spe-
cies could not be traced within the investigated material and
could possibly represent an erroneous identification of
Aequipecten macrotis. A candidate for Oppenheimopecten
aduncus is a small, abraded, single left valve found in the Sie-
ber Collection. The identification of that specimen however
on the species level appears not possible to the present author
also because the shape of the right valve remained unknown.
Previous authors (Hörnes 1867; Kautsky 1928 and Schultz
2001) illustrated only five taxa of the Grund Format-
ion. These are Crassadoma multistriata, Aequipecten macro-
tis, Macrochlamis nodosiformis, Macrochlamis tournali an d
Pecten subarcuatus styriacus. Hence the present study pro-
vides the most comprehensive documentation of the Grund
Formation pectinids hitherto.
As mentioned above one of the most important species level
revisions was made by regarding the specimens previously
identified with Hinnites brussoni leufroyi as the typical Hin-
nites crispus. The former taxon, characterizing the Late Egg-
enburgian successions (cf. Mandic & Steininger 2003) is
completely external covered with shagreen microsculpture.
Hinnitus crispus with macrosculpture almost identical with
Hinnites brussoni lacks completely the shagreen microsculp-
ture (cf. Roger 1939; Waller 1993). As even the most careful
reinvestigation of the Grund specimens, could not prove any
trace of that microsculpture on ears or disc exterior the previous
identifications by Kautsky (1928), Sieber (1947, 1949) and
Schultz (2001) were rejected and emended to Hinnites crispus.
Following the results of current reinvestigation of the type
material of Roger (1939), Almera & Bofill (1897), Kittl
(1887) and Schultz (2001) by the present author (see also
Mandic & Harzhauser 2003) Chlamys trilirata replaces
Aequipecten bryozodermis of Schultz (2001) as being found
to represent its senior synonym. The latter author already rec-
ognized the Middle Miocene specimens from the Lower Bad-
enian Gaindorf Formation (Lower Austria) and Florian For-
mation (Styria) as unrelated to the Eggenburgian Chlamys
jakloweciana (Kittl). The species is now additionally docu-
mented from the Grund Formation as well.
Following detailed reinvestigations by Studencka (1986)
the present study contradicts Schultz (2001) and reunites
Aequipecten flavus and Aequipecten malvinae. The former au-
thor showed clearly that those taxa represent only growth
stages of one and the same taxon. This is underpinned by the
fact that their type specimens (Dubois 1831) derived from one
and the same locality. Indeed the findings from the Grund
Formation representing the Aequipecten flavus morphology
are all small and represent maximally young adult growth
Beside species level revisions, the present study proposes
several genus-level emendations (see Systematic paleontolo-
gy chapter for details). Hence it neglects the phyletic relation-
ship between Costellamussiopecten and Amussiopecten an d
elevates the former subgenus of Bongrain et al. (1994) to the
genus level. Moreover the previous generic classification of
Amusium cristatum (e.g. Freneix et al. 1982; Demarcq 1990;
Studencka 1999; Schultz 2001) is rejected and its classifica-
tion with Costellamussiopecten is proposed. This is based on
phyletic as well as on morphological evidence. Thus Amusium
with its type species A. pleuronectes inhabiting Pliocene to
Recent Indo-Pacific (e.g. Eames & Cox 1956) do not posses
the scaly ornament of the dorsal ears margin. In contrast the
latter is not only typically developed in the European Pliocene
Fig. 3. Illustration shows the pectinid assemblage of the Grund Formation clearly demonstrating its Early Badenian age. The left side of the
table provides a correlation of standard geochronology, magnetostratigraphy, biostratigraphy (European Mammal, (Sub)tropical Planktonic
Foraminiferal and Calcareous Nannofossil Faunal Zones) and sequence stratigraphy (European Basins) with the regional Central Para-
tethys chronostratigraphic and biostratigraphic divisions (modified after Mandic et al. 2002). The right side shows stratigraphic distributions
of pectinid species documented by the present study from the Grund Formation. The dashed line points to an uncertain, insufficiently docu-
mented literature record. The photograph of a single left valve is provided for each species. Scale bars mark the 1 cm distances.
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 133
Amusium cristatum, but also in the whole Oligocene to Mi-
ocene lineage preceding the later species including the Grund
Formation contributor Amusium cristatum badense. Their
similarity in disc morphology is regarded as a consequence of
a parallel evolution due to best possible shape adaptation to
similarly preferred swimming behaviour and is categorized as
a striking example of homeomorphy (cf. Waller 1991). Mor-
phology with dorsal scaly ornament (crista) combined with
a three tooth pair hinge, proved to be a feature typical for the
Costellamussiopecten type species C. haueri (Michelotti).
Consequently the present author regards the European Amu-
sium as a member of the genus Costellamussiopecten.
The usage of the generic designation Macrochlamis is ex-
plicitly favoured in the present study instead of the genus
name Gigantopecten, which has found unfortunately common
usage in the newer literature (e.g. Bongrain 1988, 1992; Stu-
dencka et al. 1998; Schultz 2001). The original spelling Mac-
rochlamis by Sacco (1897a) is the correct original spelling
sensu ICZN 1985 Art. 32b because no evidence for an incor-
rect spelling is provided in the original publication according
to the same article, section c. Its emendation by Sacco (1897b)
into Macrochlamys by means of different subsequent spelling
without explanation in a following publication (Art. 33b (i))
must be discarded as being homonymic with the name already
occupied by a gastropod (comp. Hertlein 1969). The substitu-
tion of the name Macrochlamys with the new name Gigan-
topecten by Rovereto (1899) is consequently also invalid be-
cause the next available name is already provided with
Macrochlamis. The principle of one-letter difference in ge-
nus-group names (Art. 56b) provides the final argument for
the validity of the latter name.
Oopecten solarium was currently classified with Flabel-
lipecten (e.g. Bongrain 1986; Schultz 2001), which contra-
dicted the result of Roger (1939). The latter author investigat-
ed the complete European Neogene pectinid record and
concluded its closest relationship with the Oopecten rotunda-
tus (Lamarck) the type species of Oopecten (Sacco 1897b).
The present author follows the latter result and refers to the
extremely laterally elongated ears, but also to the gigantic size
that this species can attain (e.g. Kautsky 1928), a feature coin-
ciding with other related taxa like for example the Early Egg-
enburgian Oopecten gigas.
The generic emendation for Chlamys trilirata was made
by neglecting its classification with the Aequipecten proposed
by Schultz (2001). Thus Waller (1991, 1993) very clearly de-
fined Aequipecten using morphological as well as phylogenet-
ical arguments. Beside the hinge shape with a prominent later-
ally elongated resiliar tooth pair in the right valve, the
development of the commarginal sculpture with projecting
lamellae forming an inner lobe on the rib flanks are also
among the most striking features, definitely absent in
Chlamys trilirata. In contrast the latter species bears well
developed shagreen microsculpture unique for the group of
taxa closely related to the genus Chlamys in the strict sense
(cf. Waller 1993). As, however, another feature typical of
Chlamys the numerous thin radial ribs multiplying distally
by intercalation or bifurcation is absent, and another genus
level designation including forms similar to Chlamys tri-
lirata is not available the present study applies the provisional
taxonomic genus level designation. Hence the possible new
genus represented by that unusual morphology also apparent-
ly includes Chlamys jakloweciana.
The recognition of three new species (Aequipecten malvi-
nae, Hinnites cristatus, Chlamys trilirata) in the Grund For-
mation apparently improved the biostratigraphic significance
of the represented assemblage. Thus the FOD (First Occur-
rence Datum) of all three taxa is above the base of the Middle
Miocene in the Paratethys, excluding unequivocally the spec-
ulated Lower Miocene age for Grund Formation
(Fig. 3). Moreover Aequipecten malvinae with its form
Aequipecten flavus was regarded as having a high biostrati-
graphic significance in contributing the Badenian pectinid as-
semblage zone defined by Bohn-Havas et al. (1987). Its only
older record deriving from a list by Kókay (1967), which pro-
vided the authors identifications of the Karpatian molluscs
collected at Várpalota in Central Hungary is classified as
highly doubtful in the present study. No other Lower Miocene
Paratethys record of that species is additionally available.
Oopecten solarium and Macrochlamis nodosiformis are
principally unknown from the Early Miocene of the Central
Paratethys, being otherwise common in the Middle Miocene
reaching there huge dimensions (cf. Kautsky 1928). The only
possible exception is their Karpatian record in the Várpalota
Basin of Central Hungary (Kókay 1967 and 1991). Yet the
biostratigraphic re-evaluation of the Várpalota Basin deposits
formerly attributed to the Karpatian shows that at least their
upper part (the series 4. and 5. of Kókay 1967, acc. to person-
al communication to author) exactly correlates with the Grund
Horizon. Hence, as referred by Kókay (1991) they bear not
only the Badenian benthic foraminifer Borelis melo, but also
passes gradually toward the basin into pelitic-sandy sedi-
ments bearing the calcareous nannoplankton assemblage of
the NN5 Zone. From these sediments indeed the local FOD of
Aequipecten malvinae is recorded. Oopecten solarium an d
Macrochlamis nodosiformis derive, however, from the under-
lying sediments belonging to the NN4 Zone (Kókay 1991).
Their attribution to the Karpatian still remains questionable as
NN4 datum has been likely recorded from the lower part of
the Grund Formation (Æoriæ & Rögl 2004) as well as from the
Lower Badenian of the Styrian Basin (Rögl et al. 2002). Inter-
estingly, as proved by the present author in the Kókay Collec-
tion (Budapest), the oldest Central Paratethys occurrence of
Chlamys trilirata is also recorded from the latter horizon of
the I-116 well.
Oopecten solarium, Macrochlamis nodosiformis along with
Aequipecten malvinae all seem to originate in the Early Mi-
ocene of the NE Atlantic and Mediterranean (cf. Roger 1939;
Schultz 2001), migrating with the changing paleoclimate
around the Early/Middle Miocene transition into the Central
Paratethys (cf. Harzhauser et al. 2003). The Early Miocene
record of Chlamys trilirata is possibly represented by speci-
mens from the Algerian Burdigalian identified by Freneix et
al. (1974) with Chlamys jakloweciana. In contrast Hinnites
crispus is apparently absent in any Lower Miocene deposits
according to the authors knowledge (cf. Roger 1939; Waller
1993) implying unequivocally the Middle Miocene age for
the Grund Formation.
Whereas all other species are transitional throughout the
Badenian, the absence of Chlamys trilirata seems to be
proven at least from the Upper Badenian. Its only Middle
Badenian record was signalized to the author currently by
Kókay (pers. comm.) by referring to findings from explora-
tion wells for the new metroline of Budapest in Hungary. Its
presence in the Lower Lagenidae Zone is recorded throughout
Austria in the Grund-, Gaindorf- and Florian Formations.
Yet from the Austrian Upper Lagenidae Zone no records are
available. Yet, from the latter horizon of the Hungarian Várpal-
ota Basin a well preserved right valve is illustrated by Kecs-
kemeti Körmendy (1962). Interestingly, Ben Moussa & De-
marcq (1992) restricted their Chlamys jakloweciana from the
Mediterranean and NE Atlantic (most probably correspond-
ing) with Chlamys trilirata to the Langian and Serravallian.
The type locality of Chlamys trilirata is dated to the Early
Langhian (Freneix et al. 1982). The locality in the Boujan in
Provance produced a specimen rich sample originally studied
by Roger (1939) has been dated to the Helvetian and its actual
position within the Middle Miocene could not be detected
within the more modern literature.
The transitional Early to Middle Miocene character of the
represented molluscs is underlined by the common occurrence
of taxa like Pecten subarcuatus styriacus, Crassadoma multi-
striata and Aequipecten macrotis. Those species are known
already from the Late Eggenburgian to Early Ottnangian de-
posits. The FOD of Macrochlamis tournali is recorded from
the Late Ottnangian of the Várpalota Basin and from the
Karpatian of other Central Paratethyan sites. The strati-
graphically oldest record of Costellamussiopecten cristatum
badense seems to be in the Karpatian (Csepreghy-Meznerics
Whereas the pectinid assemblage of the Grund Formation
accurately indicates its Badenian age as suggested at least by
the occurrence of Aequipecten malvinae and Hinnites crispus,
a more precise bistratigraphic resolution based on the strati-
graphic ranges of recorded taxa is impoverished. Hence all
species except for Chlamys trilirata seem to be present
throughout the Badenian (Fig. 3). Nevertheless the above dis-
cussed range of Chlamys trilirata means that at least the
Late Badenian age of the Grund Formation can be accurately
Despite that, the absence of some characteristic Badenian
taxa, might be suspected as highly indicative for its lowermost
Badenian position (cf. Kautsky 1928). Hence the species typi-
cally present in the younger sediments of the Vienna Basin
such as the Upper Lagenidae Zone Baden-Sooß and Vösen-
dorf like Costellamussiopecten spinulosus and Aequipect-
en elegans are definitely absent in the Grund Formation. Ap-
parently they are also missing in the same level of the
Niederleis Basin. Studencka (1999), however, argued that the
introduction of the Badenian pectinids including Aequipecten
elegans appears to be diachronous in the Central Paratethys.
Hence whereas Aequipecten elegans is very common in the
Lower Badenian of the Pannonian Basin System allowing
Bon Havas et al. (1987) to define there the regional Chlamys
elegansPecten revolutus Subzone it is absent in the same
horizon of the Alpine-Carpathian Foredeep. For the Pannon-
ian basinal system, its record from the Lower Lagenidae Zone
of Costei in Romania is referred by Nicorici (1977) and Stu-
dencka et al. (1998). From the same locality Costellamussio-
pecten spinulosus attenuatus a form closely related with C.
spinulosus spinulosus and likely absent from the Grund For-
mation is also available. Therefore at least the absence of
the former taxon might be controlled by the paleoenvironmen-
An updating and revision of the genus and species level
taxa discarded several polyphyletic taxa and allowed a more
accurate stratigraphic and paleobiogeographical correlation
with occurrences from the Paratethys, NE Atlantic, Mediterra-
nean, and Indian Ocean. Taxonomic revision brought several
important genus-level and species level emendations. Hence
the European Amusium cristatum lineage is for the first time
classified with Costellamussiopecten. This former subgenus
of Amussiopecten has been regarded as phylogenetically unre-
lated to it and in consequence elevated to the genus level.
Macrochlamis has been regarded as a valid name, the genus-
level designation Gigantopecten was recognized as its senior
synonym. Probably a still unavailable genus-name for a
shagreen bearing Chlamys trilirata and probably also Early
Miocene Chlamys jakloweciana species related with the
Chlamys in the strict sense are provisorily termed as
Chlamys. The species level revisions include discovery of
Hinnites crispus based on previously erroneously identified
specimens. Moreover the new material brought the first evi-
dence of the presence of Ae. malvinae and Ch. trilirata in the
Grund Formation. Ae. malvinae is thereby considered to be the
senior synonym and the adult growth stage of Ae. flavus.
Based on reviewed taxonomy, this study proves that the
pectinid bivalves as an important tool in solving the regional
biostratigraphical problems. Hence, at least for the taxonomi-
cally and stratigraphically secured record from Austria, a reso-
lution corresponding to the regional foraminiferal ecozone
system has been detected. Thus the 10 species from the Grund
Formation allow an accurate regional correlation with the
Lower Lagenidae Zone and the lower part of the Lower Bade-
nian based on the absence of taxa such as Aequipecten ele-
gans and Costellamussiopecten spinulosus, which are com-
mon and typical species of the Vienna Basin Upper Lagenidae
Zone. The correlation on the Central Paratethys level approxi-
mates the resolution of the calcareous nannoplankton imply-
ing its Early Badenian and possible Middle Badenian age.
Class Bivalvia Linné, 1758
Order Ostreoida Férussac, 1822 [emend. Waller, 1978]
Family Pectinidae Wilkes, 1810 [emend. Waller, 1978]
Remark: The taxonomy of the suprageneric level is left
out in consequence that the emendation of the subfamily level
taxonomy by Waller (1993) did not achieve a clear mono-
phyletic division on that taxonomic level (Beu 1996; Waller,
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 135
pers. com. 2001; the present authors unpublished results).
The tribal division on the other hand, introduced by Waller
(1991, 1993), for the main recent taxa is left out as being ap-
parently still under construction.
Except for revisions, the species level taxonomy applied in
the present study follows Schultz (2001). As the latter author
provided extended synonymy lists for each of those taxa, the
synonymy lists of the present study refer primarily to that
Genus Chlamys (Bolten) Röding, 1798
[emend. Waller, 1993]
Type species: By Herrmannsen (1846), Pecten islandi-
cus Müller, 1776, Pleistocene to Recent, N-Atlantic, within
the cold pulses also in the Mediterranean (Roger 1939; Waller
Chlamys trilirata (Almera et Bofill, 1897)
1897 Pecten triliratus Almera et Bofill Almera et Bofill, p. 401, pl.
4, Figs. 6 and 6a
1897 Pecten bryozodermis Almera et Bofill Almera et Bofill, p. 403,
pl. 4, Figs. 13, 13a and 13b
1900 Pecten jaklowecianus Kittl Bauer, p. 46, pl. 2, Figs. 1922
1928 Chlamys jakloweciana Kittl Kautsky, p. 255 (pars)
1939 Chlamys jakloweciana Kittl Roger, p. 163 (pars), pl. 22, Figs.
910 and 1618, pl. 24, fig. 7, pl. 26, Figs. 1819
1962 Chlamys jakloweciana Kittl Kecskemeti Körmendy, p. 218, pl.
23, Figs. 12 and 17
2001 Aequipecten bryozodermis (Almera et Bofill, 1897) Schultz, p.
187, pl. 17, Fig. 8
2003 Chlamys trilirata (Almera et Bofill, 1897) Mandic et Harz-
hauser, p. 101, pl. 2, Fig. 8, pl. 3, Figs. 13
Type: Roger (1939) referred the single right valve, origi-
nally reproduced as a drawing by Almera & Bofill (1897),
with the height of 26.0 mm and the anterodorsal part broken
for the holotype. It is housed in the Type Collection of the De-
partment of Earth Sciences, Claude Bernard University Lyon
(Identification No.: 29.088). Roger (1939, Pl. 22, Fig. 16) pro-
vided its photograph. Type locality, according to Almera
(1896) and the identification card of the holotype, are expo-
sures along the railway trench from E Ermita de Bará to San
Vicente (Vendrell) in Catalonia (Tarragona, NE Spain). The
succession of marine marls, sands and detritic limestones ac-
cording to planktonic foraminiferal data of Freneix et al.
(1982) is Langhian (early Middle Miocene) in age.
Material: 2 right valves from Grund (Coll. NHMlv), 2
right and 3 left valves from E Grund (Coll. DPUV2). Addi-
tionally studied material is from the Lower Badenian of the
Gaindorf Formation (Alpine-Carpathian Foredeep in Lower
Austria) from Mühlbach, investigated by Mandic &
Harzhauser (2003), from Gaindorf investigated and identified
by Kautsky (1928) and Roger (1939) as Chlamys jakloweci-
ana (Coll. NHMs and NHMls) and from Goggendorf (Coll.
NHMls; cf. Mandic & Harzhauser 2003). The holotype and
the type specimen of Pecten bryozodermis Almera et Bofill
from the Langhian of the Bara region in NE Spain and numer-
ous other specimens from the Middle Miocene of Boujan (NE
Bezier in S France) housed in the Type Collection of the De-
partment of Earth Sciences, Claude Bernard University Lyon
and originally classified with Chlamys jakloweciana (Kittl)
were also investigated.
Dimensions: Right valve disc height = 22 mm, disc
convexity = 3.2 mm; left valve disc height = 11.2 mm,
disc length = 10.4 mm, disc convexity = 1.9 mm, umbonal
angle = 89.5°, ears length = 8.3 mm, ears height = 3.1 mm.
Description: Chlamydoid shell is small (maximally
22 cm in length), higher than long with length/high-ratio
ranging between 0.88 and 0.93, biconvex, moderately arched,
valves are thin and outside the palial line translucent, discs
both posteroventrally elongated with rounded posterior and
ventral margins and pointed anterior and dorsal margins, um-
bonal angles ranging between 83 and 90°. The right valve
with 1819 ribs, the left one with 17 ribs; on both valves ribs
moderately prominent, wider than interspaces convex to sub-
triangular, initiated non-synchronously (the subsequent ones
by intercalation) between shell height of 0.7 and 2 mm. At
shell height of about 10 mm a single riblet can intercalate the
interspaces of the lateral region, at shell height of about
20 mm three riblets on rib tops and one in each interspace be-
come initiated on the central disc region, the riblets can devel-
op coarse scales on tops. Ears large, pointed and about two
times longer anteriorly, truncated posteriorly, up to 5 riblets
can sculpture each one. Active ctenolium with fine denticles,
and deep byssal synus present. Exterior typically sculptured
by shagreen microsculpture present on the whole shell or re-
duced to its parts as in morphs with transversal bars developed
on rib tops. The smooth parts bear commonly Camptonectes-
type microsculpture. Interior with internal ribs lining the inter-
paces and missing the rib carinae. Hinge region very narrow
with reduced dentition and shallow, triangular resiliar pit.
Variation: The strong morphological variation typical
for this species (cf. Mandic & Harzhauser 2003) is expressed
in the prominence of ribs and riblets and in the development
of the shagreen microsculpture that can be almost completely
absent. The transversal bars on rib tops are also rarely present.
Normally the weaker radial sculpture is followed by reduced
shagreen microsculpture. The transitional character of this
variation is documented by a sample from the Middle Mi-
ocene of Boujan in S-France including more than 40 speci-
mens and housed in the Roger (1939) collection.
Remarks: As noted by Mandic & Harzhauser (2003) in-
vestigations of the type material housed at the Type Collec-
tion of the Department of Earth Sciences, Claude Bernard
University Lyon proved types of Chlamys triliratus an d
Chlamys bryozodermis (Almera et Bofill) for the left and
the right valve of the same species. Roger (1939) correctly
classified the latter specimen together with the specimens
from the Lower Badenian of the Gaindorf Formation. Yet as
he erroneously held them for the topotypes of Chlamys jak-
loweciana (Kittl) he applied the latter name to all those speci-
mens. The mistake was detected and partly revised by Schultz
(2001). Hence Chlamys jakloweciana is not only restricted to
the Early Miocene, but also differs by distinctly larger size,
more massive shell and less numerous but more prominent
ribs with a different sculptural pattern. The large left valve
from the Badenian of Sievring being a part of the Chlamys
jakloweciana original type series also differs from the Eggen-
burgian representatives (cf. Schultz 2001). According to the
arrangement and trigonal shape of ribs it appears to be related
to Chlamys justiniana (Fontannes).
Chlamys jakloweciana indicated by Kautsky (1928) from
the Karpatian (Late Burdigalian, Early Miocene) of Stetten/
Korneuburg (NE Austria) is an erroneous identification of
Aequipecten fragments previously published by Glaessner
(1926) as Pecten (Chlamys) sp. nova. The lot bearing an iden-
tical inscription written by Glaessner (pers. com. M.
Harzhauser, Vienna), but also Chlamys jakloweciana Kittl,
Dr. Kautsky det. is housed in the Locality Collection of the
Geological Department Natural History Museum Vienna.
As pointed out by Waller (1993) the shagreen microsculp-
ture is a feature unique to Chlamys-related pectinids. Thus
among other typical features like hinge and commarginal
lamellae shape, especially the presence of shagreen microscu-
lpture do not allow classification of Chlamys triliratus with
Aequipecten (cf. Waller 1991). However, its morphology also
differs importantly from the representatives of Chlamys s.s.
(cf. Waller 1993). Hence the genus is provisionally referred to
Stratigr./Geogr. range: Early (to Middle?) Badenian
of the Central Paratethys and Middle (to Late?) Miocene of
the Mediterranean to NE Atlantic.
Its proper Central Paratethys documentation is restricted to
the Lower Lagenidae Zone of the Gaindorf Formation, Grund
Formation and Florian Formation in Styria (Bauer 1900) and
Upper Lagenidae Zone of the Várpalota Basin (Kecskemeti
Körmendy 1962). Other records, referring to Chlamys jak-
loweciana come from different Early Badenian localities in
Romania and Bulgaria (Studencka et al. 1998). According to
personal communication with Barbara Studencka (2003) the
latter authors based their designation on a specimen from
Costei illustrated by Nicorici (1977, pl. 42, Figs. 2a2e), rep-
resenting however a different species related to Manupecten.
Its presence in the Middle Badenian of Budapest is currently
signalized to author by Kókay (personal communication).
Fragments of Chlamys jakloweciana from the Badenian of
Samsonhaza, illustrated by Csepreghy-Meznerics (1960, pl.
16, Figs. 67) are not related to Chlamys trilirata and there-
fore the reference of its presence in the Karpatian of
Püspökhatvan and Ottnangian of Budapest-Cinkota in the
same study must be doubted.
The extraordinary rich collection from Boujan in Southern
France originally documented by Roger (1939) is dated to the
Helvetien and can be referred to the Middle Miocene. The
type locality is of Langhian age (Freneix et al. 1982). Ser-
ravallian and Tortonian occurrence of Chlamys jakloweciana
is mentioned only by Dermitzakis & Georgiades-Dikeoulia
(1984), but without material evidence it is doubted in the
Genus Hinnites Defrance, 1821
Type species: Ostrea crispa Brocchi, 1814
Hinnites crispus (Brocchi, 1814)
1867 Hinnites defrancei Micht. Hörnes, p. 423 (pars), pl. 2, Figs.
1928 Hinnites leufroyi Serr. Kautsky, p. 264
1947 Hinnites leufroyi Serr. Sieber, p. 112
1939 Chlamys crispa (Brocchi, 1829) Roger, p. 172, pl. 23, Figs. 11,
12, pl. 24, Fig. 6, pl. 25, Figs. 1, 4, pl. 28, Fig. 2
2001 Hinnites brussoni leufroyi (de Serres, 1829) Schultz, p. 221
(pars), pl. 20, Figs. 2ab, 4ab
Type: Rossi-Ronchetti (19511957) referred to an articu-
lated specimen illustrated by Brocchi (1814) and housed in
the Natural History Museum in Milan (Brocchi collection;
No. 460) for the holotype. It derived from Asti in NW Italy
(cf. Roger 1939). The age of that occurrence originally regard-
ed as Piacentino is Piacenzian (early Late Pliocene).
Material: Specimens from Windpassing (Coll. NHMls)
and Grund (Coll. NHMs).
Dimensions: Right valve disc height = 46.2 mm,
disc length = 42 mm, disc convexity = 4 mm, umbonal angle
= 102°, ears length = 26.7 mm, ears height = 10.2 mm; left
valve disc height = 62.3 mm, disc length = 57 mm, disc
convexity = 13 mm, umbonal angle = 106°, ears length =
34.6 mm, ears height = 15 mm.
Remark: The studied specimens were previously errone-
ously identified with Hinnites brussoni (compare Schultz
2001) yet also the most careful reinvestigation could not de-
tect any traces of shagreen microsculpture which is its main
distinctive feature toward Hinnites crispus (compare Roger
1939; Waller 1993). The apparent difference between former
species is indeed only the absence of shagreen microsculpture
in Hinnites crispus (cf. Roger 1939). Indeed both species are
represented in the Middle Miocene of Paratethys. Hence the
specimens from Grund identified by Schultz (2001) as Hin-
nites leufreyi and Hinnites defrancei, bear no shagreen mi-
crosculpture and consequently represent Hinnites crispus
morphology. In contrast the specimen illustrated by Nicorici
(1977, Pl. 44, Fig. 1a1c) and identified as Chlamys brussoni
defrancei, from the Badenian of SW Romania show shagreen
microsculptural pattern and consequently represent typical
Hinnites brussoni. With minor exceptions the present author
principally follows revisions introduced by Waller (1991 and
1993). One such exception is the status of Hinnites crispus
which is in the present study considered as a direct descendant
of Hinnites brussoni. This interpretation supported by related
shell morphologies as well as by their stratigraphic ranges,
stays in discordance with the inference by Waller (1993) ad-
vocating the branching of Hinnites crispus off a Laevichlamys
Stratigr./Geogr. range: Middle Miocene to Plio-
cene, ?Recent/Central Paratethys, Mediterranean. In the Cen-
tral Paratethys it is restricted to the Badenian, in the Mediter-
ranean and E Atlantic regions it is typically present from the
Middle Miocene to Pliocene (Roger 1939). Waller (1993) re-
fers the living Hinnites from the tropical West Africa offshore
also to Hinnites crispus. It is possible that the latter region
represents the current refugium of this termophylic taxon.
Genus Crassadoma Bernard, 1986
[emend. Waller, 1993]
Type species: By original designation (Bernard, 1986),
Lima gigantea Gray; Miocene to Recent of the eastern Pacific
(Baja California to Alaska).
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 137
Fig. 4. Chlamys, Hinnites and Crassadoma from the Grund Formation. 15 Chlamys trilirata (Almera et Bofill); locality: E Grund;
collection: DPUV2 (compare text for abbreviations). 1 anterodorsally fragmented right valve showing distal introduction of secondary
radial sculpture. 2 left valve with missing anterior ear. 3 right valve with missing ears and proximal disc region. 4 a complete right
valve showing intensive shagreen microsculpture. 5 right valve with marginally fragmented disc showing almost completely reduced
shagreen microsculpture and well developed commarginal bars on rib tops characterizing the early growth stage. 67 Hinnites crispus
(Brocchi). 6 a complete right valve; locality: Windpassing; collection: NHMls. 7 a complete left valve; locality: Grund; collection:
NHMs. 89 Crassadoma multistriata (Poli) locality: E Grund; collection: DPUV2. 8 right valve with fragmented ventral margin.
9 left valve with fragmented ventral margin. (The single scale bars (1 cm) refer to specimens 6 and 7 and to specimen groups 14 and 8
9. The letters imply the view: (a) exterior, (b) interior, (c) posterior and (d) dorsal.)
Crassadoma multistriata Poli (1795)
1993 Crassadoma multistriata (Poli, 1795) Waller, p. 212, Figs.
2001 Crassadoma? multistriata s.l. (Poli, 1795) Schultz, p. 176, pl.
16, Figs. 6, 912, pl. 17, Fig. 14
Type: Waller (1993) defined the original figure of Poli
(1795, Fig. 14) as lectotype, although the original specimen,
from the Holocene of Sicily (Italy) got lost. In consequence
the author defined additionally three syntypes (Coll. NHM
Material: Numerous specimens from the field ditch E
Grund (Coll. DPUV2) and from the excavation Grund 1998
(Coll. DPUV1 and NHMlv).
Dimensions: Right valve disc height = 27 mm, disc
length = 22.2 mm, disc convexity = 3.6 mm, umbonal an-
gle = 81°, ears length = 16.1 mm, ears height = 6.5 mm; left
valve disc height = 28.9 mm, disc length = 23.8 mm, disc
convexity = 5.2 mm, umbonal angle = 82.5°, ears length =
16 mm, ears height = 9 mm.
Remark: It can be easily distinguished from similar spe-
cies like Chlamys costai (Sowerby in Smith) or Chlamys jus-
tiniana (Fonntannes) by completely lacking the shagreen mi-
crosculpture. Chlamys varia (Linné) differs by its fixed
number of ribs early in the ontogeny.
Stratigr./Geogr. range: Late Eggenburgian to Bade-
nian of the Central Paratethys, Burdigalian Upper Marine Mo-
lasse of the Western Paratethys and Sakaraulian to Tarkhanian
and Konkian of the Eastern Paratethys; Priabonian?, Burdiga-
lian to Recent of the Mediterranean, E Atlantic and Indian
Crassadoma multistriata inhabited at least since Burdiga-
lian the regions between the Atlantic and the Indian Ocean,
although its oldest reference is dated to the Priabonian (cf.
Waller 1993). Its Central Paratethyan record apparently be-
gins with the Late Eggenburgian when it immediately be-
comes common and characteristic within the shallow water
successions (cf. Mandic & Steininger 2003).
Genus Aequipecten Fischer, 1886
Type species: By original designation (Fischer 1886)
Ostrea opercularis Linnaeus, 1758 Recent eastern Atlantic
Aequipecten macrotis (Sowerby in Smith, 1847)
1949 Chlamys macrotis Sow. Sieber, p. 112
2001 Aequipecten macrotis macrotis (Sowerby, 1847) Schultz, p.
192, pl. 18, Fig. 2
Type: The holotype was originally not designated, yet
Ètyroký (1969) regarded the left valve illustrated by Sowerby
in Smith (1847, pl. 17, Fig. 15) as the holotype by monotypy.
The specimen was collected at Adica, S Lisbon or Piedade re-
gion in Portugal. Apparently the type series included speci-
mens from both localities and according to ICZN 1985, Art.
74b, Ètyroký (1969) must be deemed to have designated the
lectotype. Veiga Ferreira (1969) referred the occurrence at
Adica to the Tortonian.
Material: Numerous juvenile to adult specimens from
the field ditch E Grund (Coll. DPVUV2) from the Grund ex-
cavation 1998 (Coll. DPUV1 and NHMlv) and from Immen-
dorf (Coll. NHMls).
Dimensions: Right valve disc height = 30.1 mm, disc
length = 30.1 mm, disc convexity = 6.5 mm, umbonal an-
gle = 82.5°, ears length = 21.2 mm, ears height = 8.5 mm; left
valve disc height = 25.8 mm, disc length = 27 mm, disc
convexity = 6 mm, umbonal angle = 98°, ears length = 17 mm,
ears height = 7 mm.
Remark: With 18 to 24 ribs this characteristic species has
significantly fewer ribs than Aequipecten malvinae.
Aequipecten macrotis is typically represented in the Early
Badenian sediments of Grund and Niederleis. At Niederleis,
it is the most typical molluscan species of the maerl facies
(Mandic et al. 2002). In the latter locality it has been charac-
teristically found is a marl facies. In the Paratethys it is
present in the Burdigalian Upper Marine Molasse of the West-
ern Paratethys. In the Central Paratethys the oldest findings
are from the late Ottnangian of Bantapuszta in Hungary (see
Stratigr./Geogr. range: Late Ottnangian to Bade-
nian of the Central Paratethys. Miocene of the Mediterranean
to NE Atlantic.
Aequipecten malvinae (Dubois, 1831)
1986 Chlamys (Aequipecten) malvinae (du Bois de Montpereux,
1831) Studencka, 35, pl. 4, Figs. 4, 7 and 10ab
2001 Aequipecten flavus (Dubois, 1831) Schultz, p. 191, pl. 18,
2001 Aequipecten malvinae (Dubois de Montpereux, 1831) Schultz,
p. 197, pl. 18, Fig. 4 and 5
Type: Specimens illustrated by Dubois de Montpereux
(1831, Pl. 8, Figs. 2 and 3) are from Shushkivtsky (former
Szuszkowce, spelling by Dubois as Szuskowce) in the Ter-
nopol region, NW Ukraine. Occurrence is dated to the Late
Badenian (Muratov & Nevesskaja 1986; Studencka & Popov
1996). Aequipecten flavus (Dubois) and Aequipecten diapha-
nus (Dubois) types are from the same locality.
Material: Specimens from the field ditch E Grund (Coll.
DPUV2) and from the Grund excavation 1998 (Coll. NHMlv).
Dimensions: Right valve disc height = 15.6 mm, disc
length = 15 mm, disc convexity = 3 mm, umbonal angle = 95°,
ears height = 3.5 mm; left valve disc height = 22.1 mm, disc
length = 21.6 mm, disc convexity = 4.9 mm, umbonal angle =
95°, ears length = 13.8 mm, ears height = 5 mm.
Remark: The reunion of Aequipecten flavus an d
Aequipecten malvinae follows a great deal of evidence pro-
vided by Studencka (1986). The decision was moreover facili-
tated by the fact that the type series of the latter species de-
rived from the same locality. The extremely detailed
reinvestigation of numerous specimens from the type region
could show that Ae. flavus represent apparently the young
adult stage of Ae. malvinae. This correlates with the fact that
specimens derived from the Grund Formation, varying by the
rib number between 37 and 43, hence corresponding with Ae.
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 139
Fig. 5. Aequipecten, Costellamussiopecten and Oopecten from the Grund Formation. 12 Aequipecten macrotis (Sowerby in
Smith) locality: Immendorf; collection: NHMls (compare text for abbreviations). 1 left valve with missing anterodorsal ear por-
tion. 2 a complete right valve. 34 Aequipecten malvinae (Dubois) locality: Grund; collection: DPUV2. 3 a well preserved
left valve. 4 right valve missing the anterior ear. 57 Costellamussiopecten cristatus badense (Fontannes) locality: Windpass-
ing; collection: NHMls. 5 right valve with fragmented ventral disc portion. 6 right valve with fragmented postero- and anterodor-
sal margins. 7 left valve with abraded margins. 8 Oopecten solarium (Lamarck) locality: Windpassing; collection: NHMls. 8
a single left valve. (The single scale bars (1 cm) refer to specimens 5 and 7 and to specimen groups 12, 24 and 67. The letters imply
the view: (a) exterior, (b) interior, (c) posterior and (d) dorsal.)
flavus are all smaller than 22 mm in height. According to
Schultz (2001) Ae. malvinae has, besides better-developed
secondary sculpture, also the lowered rib number (2730 with
23 lateral ones).
Stratigr./Geogr. range: Badenian of the Central
Paratethys, Tarkhanian and Konkian of the Eastern Para-
tethys, Burdigalian to Tortonian of the Mediterranean to NE
Specimen from the Late Ottnangian (Bohn-Havas et al.
1985) of Bantapuszta in the Central Hungarian Várpalota Ba-
sin identified by Kókay in Steininger et al. (1973) with
Aequipecten malvinae apparently has a smaller number of ribs
and represents another Aequipecten related species. As dis-
cussed above, the Karpatian occurrence from the same basin
(Kókay 1967) is reconsidered for being Early Badenian in
age. This species remains in consequence unknown from lay-
ers older than Badenian. Subsequently also Aequipecten fla-
vus is definitely restricted to the Badenian (e.g. Bohn-Havas
et al. 1987).
Genus Costellamussiopecten Bongrain, Cahuzac et Freneix,
Type species: By original designation (Bongrain et al.
1994) Pecten haueri Michelotti from the Miocene of the NE-
Atlantic, Mediterranean and Paratethys.
Diagnosis: Following Bongrain et al. (1994), a list of
features characterizing the genus can be given: (1) right hinge
with 3 tooth pairs, whereby the dorsal tooth pair dominates,
(2) spiny to scaly radial secondary sculpture at least early in
the ontogeny, (3) dentition of the right dorsal edge (= crista),
(4) presence of Camptonectes microsculpture, (5) smooth pre-
radial stage in the left valve and (6) distinct aragonite layer in-
tercalated within the umbonal region. A diagnosis emendation
given herein pertains to the presence of aequipectinoid in ad-
dition to amussiopectinoid disc shapes in the genus.
Remarks: Costellamussiopecten included according to
the original designation by Bongrain et al. (1994) a group of
secondarily, scaly sculptured species, previously classified
with Aequipecten and related to C. haueri. It has been defined
as a subgenus of Amussiopecten based on its apparent rela-
tionship to C. baranensis (a Middle Miocene NE-Spain spe-
cies closely related to C. cristatus badense), classified errone-
ously with the latter genus. Yet, Amussiopecten, along with
Amussium never develop the scaly secondary sculpture, at
best exampled by the dorsal crista and can therefore not be re-
lated to Costellamussiopecten. Representing apparently an in-
dependent taxonomic unit, it has been elevated by the present
study onto the genus level.
The ongoing investigations by the present author (Mandic,
in prep.) shows that the genus has Oligocene to Pliocene
range and NE-Atlantic, Mediterranean and Paratethys distri-
bution. The following species recorded in the Paratethys are
tentatively considered to be related to the genus: Costellamus-
siopecten haueri (Michelotti, 1847), C. koheni (Fuchs, 1876),
C. spinulosus (Münster in Goldfuss, 18341840), C. pasinii
(Meneghini, 1857), C. cristatus badense (Fontannes, 1882),
C. deletus (Michelotti, 1861), C. telegdirothi (Csepreghy-
Meznerics, 1960), C. northamptoni (Michelotti, 1839). Addi-
tional species possibly included are Costellamussiopecten?
schreteri (Noszky, 1936), C.? oligosquamosus (Sacco, 1897),
C.? perrugosus (Sacco, 1897), and C.? martelli (Ugolini,
Costellamussiopecten cristatus badense (Fontannes, 1882)
1867 Pecten cristatus Bronn. Hörnes (pars.), p. 419, pl. 66, Figs. 1ad
1928 Amussium cristatum Bronn. var. badensis Font. Kautsky, p. 253
1947 Amussium cristatum Bronn. var. badensis Font. Sieber, p. 158
2001 Amusium cristatum badense (Fontannes, 1882) Schultz, p.
157, pl. 15, Figs. 13ab
Type: The type is the articulated specimen illustrated by
Hörnes (1867) housed in the NHMs collection. It derived
from marls at Möllersdorf in Lower Austria, dated as Upper
Lagenidae Zone (Early Badenian).
Material: Specimens from Windpassing (Coll. NHMls).
Dimensions: Right valve disc height = 28 mm, disc
length = 29.8 mm, disc convexity = 3.1 mm, umbonal an-
gle = 119.5°, ears length = 15 mm, ears height = 6 mm; left
valve disc height = 33.8 mm, disc length = 33 mm, disc
convexity = 3 mm, umbonal angle = 121°, ears length = 19 mm,
ears height = 5.5 mm.
Remarks: This taxon belongs to a lineage of amussiopec-
tinoid shaped Costellamussium, which originated in the Oli-
gocene, possibly from Costellamussiopecten deletus. Its old-
est representative is C. pasini (Meneghini). The lineage
becomes extinct in the Pliocene. Its last representative was the
European Amusium Amusium cristatum (Bronn.). The
morphologically homologue AmussiopectenAmusium lin-
eage originated likely in the Oligocene, but as mentioned above
never developed the scaly ornament and the dorsal crista.
Costellamussiopecten cristatus badense is morphologically
closely related to its Pliocene successor, hence some authors
tend to regard them as synonymous (e.g. Studencka et al.
1998). In contrast Schultz (2001) following Kautsky (1928)
consider the difference as allowing the subspecies level divi-
sion. Principally Costellamussiopecten cristatus has mostly
completely suppressed ribs of the left umbonal region being
moreover dorso-ventrally elongated.
Stratigr./Geogr. range: Karpatian to Badenian of
the Central Paratethys, Burdigalian to Messinian of the Medi-
terranean to NE Atlantic (Schultz 2001).
Genus Oopecten Sacco, 1897a
Type species: By original designation (Sacco 1897a)
Pecten rotundatus Lam., Burdigalian of southeastern France.
Oopecten solarium (Lamarck, 1819)
1867 Pecten besseri Andrz. Hörnes, 404 (pars), pl. 62, Figs.12, (?: pl.
63, Figs. 15)
1928 Pecten (Amussiopecten) solarium Lamarck Kautsky, p. 250
1947 Pecten (Amussiopecten) solarium Lam. Sieber, p. 158
2001 Flabellipecten solarium (Lamarck, 1819) Schultz, p. 236 (pars)
Type: The holotype is not designated. The type series of
Lamarck (1819) derivated from the Doue environ in the Loire
Basin (NW France) (cf. Dollfuss & Dautzenberg 19021920).
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 141
According to Bongrain (1988) those sediments bear land
mammal association of the MN9 European Faunal Zone cor-
relating with the Tortonian (cf. Steininger et al. 1996).
Material: Two left valves from Windpassing (NHMW l.c.).
D i m e n s i o n s : Left valve disc height = 54 mm, disc
length = 58 mm, disc convexity = 7.5, umbonal angle = 125°,
ears length = 38 mm, ears height = 10.5 mm.
Description: The left valve is of moderate size, with
disc rounded for most of the outline except for slightly con-
cave postero- and anterodorsal margins, very slightly elongat-
ed posteroventrally, weakly convex in dorsoventral cross-sec-
tion, with umbonal angle attaining about 125°, with flattened
proximal region, lateral disc areas moderately arched, broadly
triangular in left view, similarly broad but anterior one dis-
tinctly shorter than the posterior one, anterior lateral area with
3 ribs on its posterior half, the middle one marks the maxi-
mum convexity axis, the anterior half smooth; the posterior
disc area with 6 smooth ribs on its anterior two thirds, the 3
one marks the axis of maximum convexity, the posterior third
is smooth; disc with 14 first order, prominent, flat-topped ribs,
trapezoidal in cross-section, as broad as flat-bottomed inter-
spaces, the latter with extremely weak mediate riblet, disc sur-
face covered with slightly eroded commarginal lamellae being
proximally loosely and distally densely arranged. Preradial
stage about 2 mm in height. Ears very long but low, with
straight dorsal and convex lateral margins, posterior ear larger
than the anterior ear, with the summed length about three
quarters of the disc length, the anterior one low, the poster one
moderately high, posterior ear flattened anterior ear radially
concave, with 4 dorsally prominent riblets, the contacts with
the disc low but distinct. Interiorly 15 and a half pairs of
prominent interior rib carinae. Cardinal crura with the paired
posterior denticle. Hinge with 3 to 4 tooth pairs.
Remark: The present study provides the first illustration
of an Oopecten solarium from the Grund Formation.
This species is much confused in the literature although
showing apparent distinctive features. Indeed the combination
of a rounded disc outline, extremely wide umbonal angle,
very long but low ears, numerous, typically flattened, but
prominent ribs, with single weak, intercalated riblet in inter-
spaces, wide marginal disc areas and symmetrical radiating
riblets on ears of the left valve reaching in the adult stage gi-
gantic dimensions allow an easy species identification.
Initially Hörnes (1867) classified Oopecten solarium speci-
mens together with specimens of Pecten subarcuatus styria-
cus and Pecten besseri under the latter name. Deperet & Ro-
man (19021912) following Ugolini (1908) regarded the
specimens illustrated by Hörnes (1867) as independent spe-
cies and denoted it with Pecten (Amussiopecten) incrassatus
Partsch. The latter name is however a nomen nudum of
Partsch in Hörnes (1848) that have been made available by
bibliographic reference (cf. ICZN 1985, Art. 11,d,ii and Art.
12) by Hörnes (1867). Actually the latter author included the
Partsch designation into his Pecten besseri synonymy. Doll-
fus & Dautzenberg (19021920), referring neither to Ugolini
(1908) nor to Deperet & Roman (1912), included the speci-
mens illustrated as Pecten besseri by Hörnes (1867) into Pect-
en (Oopecten) solarium. Finally Roger (1939), following
Kautsky (1928) and Friedberg (1936), reunited all those refer-
ences into Chlamys solarium (Lamarck).
Kojumdgieva (1960), Csepreghy-Meznerics (1960), Nicori-
ci (1977) and Steininger et al. (1978) document typical Cen-
tral Paratethys morphologies of Oopecten solarium. In contrast,
beside characteristic specimens illustrated by vagrovský
(1981, pl. 7 and 9) from the Upper Badenian of Devínska
Nová Ves, Slovakia, an articulated valve from his pl. 8 with
lowered number of prominent, rounded ribs and low umbonal
angle, represents more likely a Pecten or a Flabellipecten. The
same holds true for a right and a left valve documented by
Schultz (2001, pl. 38, Fig. 1 and pl. 40, Figs. 1 and 2). Hence
the left valve with marginal areas sculptured by prominent
first order ribs and ears missing the radiating riblets is appar-
ently not an Oopecten solarium. Yet, conspicuously, these
specimens are referred as Hörnes (1867, pl. 63, Figs. 4 to 5;
sic?: cf. Figs. 1 to 5, my rem.) originals, implying that the sim-
ilarity of the latter drawings with actual Oopecten solarium
must be a coincidence.
The present author follows Roger (1939) by classifying the
species with Oopecten, Bongrain (1988) and Schultz (2001)
classified it currently with Flabellipecten, but unfortunately
failed to give the arguments.
Stratigr./Geogr. range: Common and characteristic
during the whole range of the Badenian in the Central Para-
tethys. Its possible Karpatian occurrence (Kókay 1967) is dis-
cussed elsewhere in the text (see chapter Biostratigraphic
inferences). Based on the latter record Bohn-Havas et al.
(1987) also referred to its questionable occurrence from the
Karpatian. In the Mediterranean to E Atlantic region it ranges
from the Burdigalian to the Messinian (Roger 1939).
Genus Macrochlamis Sacco, 1897a
Type species: By subsequent designation (Sacco 1897b)
Ostrea latissima Brocchi from the Pliocene of Italy. Note that
Bongrain (1988, p. 228) proposed replacement of the current
type species by Pecten ligerianus Dollfuss et Dautzenberg
from the Middle Miocene of NW-France.
Macrochlamis nodosiformis (de Serres in Pusch, 1837)
1867 Pecten latissimus Brocchi Hörnes, p. 395, pl. 56, Figs. 14, pl.
57, Figs. 14
1928 Pecten (Oopecten) latissimus Brocchi var. austriaca nov.var.
Kautsky, p. 252
1949 Pecten (Oopecten) latissimus Brocchi Sieber, p. 112
2001 Gigantopecten nodosiformis (Pusch, 1837) Schultz, p. 249, pl.
36, Fig. 3, pl. 37, Figs. 12, pl. 38, Fig. 2, pl. 39
Type: The holotype was not designated; the specimen
originally illustrated by Pusch (1837, Pl. 5, Figs. 9ac) is from
Skotniki, southern slopes of the Holy Cross Mts, Central Po-
land. The type series derived from Skotniki and Widuchowa
near Busko and Kików near Stobnica. It is Early Badenian in
age (Studencka & Studencki 1988).
Material: Specimens from Windpassing and Guntersdorf
Dimensions: Right valve disc height = 71.5 mm, disc
length = 76 mm, disc convexity = 23.5 mm, umbonal an-
gle = 102°, ears length = 48 mm, ears height = 23 mm; left
valve disc height = 119.8 mm, disc length = 137 mm, disc
convexity = 39 mm, umbonal angle = 103°, ears length =
79 mm, ears height = 32 mm.
Remark: This characteristic species resembles somewhat
the Macrochlamis holgeri (Geinitz) from the Eggenburgian
deposits of Lower Austria. The latter species, however, never
develops node series on rib tops.
The generic designation follows the argumentation of
Smith (1991) regarding Gigantopecten for the junior syn-
onym of Macrochlamis (see chapter Faunal composition
and taxonomic revisions).
Stratigr./Geogr. range: It is typical and common
throughout the Badenian (cf. Schultz 2001). Its record from
the deposits underlying the Grund Horizon of the Várpalota
Basin, discussed above in the text, is Karpatian to Early Bade-
nian in age. Its first occurrence in the Mediterranean to NE
Atlantic region is in the Late Burdigalian. The species is cor-
respondingly commonly present there in the Middle and also
in the Late Miocene. In the Late Miocene (Roger 1939) or
Pliocene (Bongrain 1992) it gets replaced by the Macrochla-
Macrochlamis tournali (de Serres, 1829)
1867 Pecten tournali Serres Hörnes, p. 398, pl. 58, Figs. 16
1928 Pecten (Oopecten) tournali de Serr. Kautsky, 252
1947 Pecten (Oopecten) tournali Serr. Sieber, p. 158
1949 Pecten tournali Serr. Sieber, p. 112
2001 Gigantopecten tournali (de Serres, 1829) Schultz, p. 254, pl.
41, Figs. 12
Type: Holotype was not designated. The accurate origin
of the left valve illustrated by Serres (1829) is unknown. The
collection is derived from the marine Tertiary of southern
France (terrains marins tertiarires du midi de la France).
Indications by Roger (1939), Bongrain (1992) and Schultz
(2001) point out that the occurrence of this species in south-
ern France could be restricted to the upper Burdigalian.
Material: Specimens from Windpassing and Immendorf
Dimensions: Right valve disc height = 29.5 mm, disc
length = 27.1 mm, disc convexity = 5.7 mm, umbonal an-
gle = 100°, ears length = 19 mm, ears height = 8.5 mm; left
valve disc height = 38 mm, disc length = 41.2 mm, disc
convexity = 7.2 mm, umbonal angle = 92°, ears length =
26.8 mm, ears height = 12 mm.
Remark: Bongrain (1992) in her proposal for Gigan-
topecten, i.e. Macrochlamis, phylogeny restricted the range of
Macrochlamis tournali to the Middle to Late Burdigalian. She
derived M. tournali from Macrochlamis ziziniae (Blancken-
horn) which is a species indeed closely related to it. Yet the
author also let M. ziziniae terminate shortly above the Burdi-
galian/Langhian boundary. Its successor, an unnamed species
termed G. sp.? arose by gradual transition already in the
Early Langhian. The latter represents a morphological stage
preceding the last stage of the transition provided by the mid-
Serravalian to Tortonian Macrochlamis albina (Teppner). The
actual FOD of Macrochlamis albina is still within the Early
and not the Middle Miocene. Hence M. albina reaches its
maximum already in the Early Badenian deriving forms larger
than 20 cm (e.g. Schultz 2001).
Stratigr./Geogr. range: Late Ottnangian to Bade-
nian of the Central Paratethys. Upper Marine Molasse of the
Western Paratethys, Burdigalian to ?Tortonian of the Mediter-
ranean to NE Alantic.
Genus Pecten Müller, 1776
Type species: By subsequent designation (Schmidt
1818), Ostrea jacobaea Linneus, Pliocene to Recent of the
Mediterranean and northeastern Atlantic.
Pecten subarcuatus styriacus Hilber, 1879
1879 Pecten styriacus Hilb. Hilber, p. 455, pl. 6, Figs. 1315
1928 Pecten subarcuatus Tournouer. var. styriaca Hilber. Kautsky,
pl. 7, Figs. 910
1947 Pecten subarcuatus Tourn. var. styriaca Hilb. Sieber, p. 158
2001 Pecten subarcuatus styriacus Hilber, 1879 Schultz, p. 271, pl.
42, Figs. 23
Type: The holotype was originally not designated. The
type series consists of at least three specimens illustrated by
Hilber (1879, pl. 6, Figs. 1315). Schultz (2001) referred the
holotype, that is indeed the potential lectotype, as being
housed in the Collection of the Joanneum Museum at Graz
(SE Austria). The type series derives from the St. Florian
(= Groß St. Florian) in Styria, from sandy marls. The sedi-
ments belong to the Florian Formation of the western Styrian
Basin and are dated correspondingly to the Grund Formation
with the regional Lower Lagenidae Zone of the Early Bade-
nian (Tollmann 1985; Schultz 2001).
Material: Numerous juvenile to adult specimens from
the field ditch E Grund (Coll. DPUV2) and excavation Grund
1998 (Coll. NHMlv) and Windpassing (Coll. NHMls).
Dimensions: Left valve disc height = 38 mm, disc
length = 42.5 mm, disc convexity = 1.5 mm, umbonal an-
gle = 113°, ears length = 27.9 mm, ears height = 11 mm; right
valve disc height = 38 mm, disc length = 43.3 mm, disc
convexity = 11 mm, umbonal angle = 94°, ears length =
23.8 mm, ears height = 9 mm.
Remark: Studencka (1986) reunited P. subarcuatus with
P. styriacus which was followed by Studencka et al. (1998).
Yet, P. subarcuatus styriacus has always flat-topped ribs, as
confirmed in numerous specimens from the Grund Formation.
P. s. subarcuatus differs in that its ribs on the left and on the
right valve are convexly rounded and P. subarcuatus fuchsi
has left ribs sub-trapezoidal and the right ribs convexly round-
ed to sub-trigonal as documented by numerous specimens
from the Late Burdigalian of the Suez region in NE Egypt
(Coll. Mandic, Natural History Museum Vienna). Indeed
these differences are minute and reflect maximally the subspe-
cies level taxonomy, but coincide apparently with the paleo-
geographical distribution of those otherwise coeval taxa.
PECTINID BIVALVES FROM THE GRUND FORMATION TAXONOMIC REVISION 143
Fig. 6. Macrochlamis from the Grund Formation. 12 Macrochlamis nodosiformis (de Serres in Pusch). 1 left valve missing the ante-
rior ear; locality: Guntersdorf; collection: NHMls (compare text for abbreviations). 2 right valve; locality: Immendorf; collection:
NHMls. 34 Macrochlamis tournali (de Serres). 3 left valve missing the portions of anterodorsal ear and the ventral disc margin; lo-
cality: Immendorf; collection: NHMls. 4 a complete right valve; locality: Windpassing; collection: NHMls. (The single scale bars (1 cm)
refer to specimens 1 and 2 and to specimen group 34. The letters imply the view: (a) exterior, (b) interior, (c) posterior and (d) dorsal.)
Fig. 7. Pecten subarcuatus styriacus Hilber from the Grund For-
mation locality: Grund; collection: DPUV2. 1 left valve
with fragmented anterodorsal ear portion. 2 a complete right
valve. (The scale bar = 1 cm. The letters imply the view: (a) exte-
rior, (b) interior and (c) posterior.)
Hence P. s. subarcuatus inhabited the northeastern Atlantic,
P. s. styriacus the Central Paratethys and P. s. fuchsi the pro-
Stratigr./Geogr. range: Restricted to the Late Egg-
enburgian to Late Badenian of the Central Paratethys (cf.
Acknowledgments: Thanks go to Peter Pervesler (University
of Vienna) and Rainhard Roetzel (Geological Survey Vienna)
for organizing the excavation campaigns at Grund which gave
rise to the present study as well as for interesting discussions
about the sedimentological and paleobiological features of the
Grund Formation. Many thanks also to reviewers Barbara
Studencka (Museum of the Earth, PAS, Warsaw) and Josef
Kókay (Budapest) for thoughtful comments improving the
manuscript. This work gained also much profit from discus-
sions and cooperation with Mathias Harzhauser, Fred Rögl
(both Natural History Museum Vienna) and Martin Zuschin
(University of Vienna). Finally thanks go to Ortwin Schultz
and Mathias Harzhauser for facilitating the best possible
working conditions in the Natural History Museum Collec-
tions in Vienna and last but not least to Abel Prieur (Claude
Bernard University of Lyon) whose generous help with the
Roger (1939) type collection, made some of important taxo-
nomic revisions possible. The present study is a partial result
of the Austrian Science Fundation (FWF) Project: Evolution
vs. Migration in Austrian Miocene Mollusc Communities
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